The Origin of Species by Means of Natural Selection

Thus, as it seems to me, the manner in which single species and whole groups of species become extinct accords well with the theory of natural selection. We need not marvel at extinction; if we must marvel, let it be at our presumption in imagining for a moment that we understand the many complex contingencies on which the existence of each species depends. If we forget for an instant that each species tends to increase inordinately, and that some check is always in action, yet seldom perceived by us, the whole economy of nature will be utterly obscured. Whenever we can precisely say why this species is more abundant in individuals than that; why this species and not another can be naturalised in a given country; then, and not until then, we may justly feel surprise why we cannot account for the extinction of any particular species or group of species.

ON THE FORMS OF LIFE CHANGING ALMOST SIMULTANEOUSLY THROUGHOUT THE WORLD.

Scarcely any palaeontological discovery is more striking than the fact that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant regions, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the organic remains in certain beds present an unmistakable resemblance to those of the Chalk. It is not that the same species are met with; for in some cases not one species is identically the same, but they belong to the same families, genera, and sections of genera, and sometimes are similarly characterised in such trifling points as mere superficial sculpture. Moreover, other forms, which are not found in the Chalk of Europe, but which occur in the formations either above or below, occur in the same order at these distant points of the world. In the several successive palaeozoic formations of Russia, Western Europe and North America, a similar parallelism in the forms of life has been observed by several authors; so it is, according to Lyell, with the European and North American tertiary deposits. Even if the few fossil species which are common to the Old and New Worlds were kept wholly out of view, the general parallelism in the successive forms of life, in the palaeozoic and tertiary stages, would still be manifest, and the several formations could be easily correlated.

These observations, however, relate to the marine inhabitants of the world: we have not sufficient data to judge whether the productions of the land and of fresh water at distant points change in the same parallel manner. We may doubt whether they have thus changed: if the Megatherium, Mylodon, Macrauchenia, and Toxodon had been brought to Europe from La Plata, without any information in regard to their geological position, no one would have suspected that they had co-existed with sea-shells all still living; but as these anomalous monsters co-existed with the Mastodon and Horse, it might at least have been inferred that they had lived during one of the later tertiary stages.

When the marine forms of life are spoken of as having changed simultaneously throughout the world, it must not be supposed that this expression relates to the same year, or even to the same century, or even that it has a very strict geological sense; for if all the marine animals now living in Europe, and all those that lived in Europe during the pleistocene period (a very remote period as measured by years, including the whole glacial epoch) were compared with those now existing in South America or in Australia, the most skilful naturalist would hardly be able to say whether the present or the pleistocene inhabitants of Europe resembled most closely those of the southern hemisphere. So, again, several highly competent observers maintain that the existing productions of the United States are more closely related to those which lived in Europe during certain late tertiary stages, than to the present inhabitants of Europe; and if this be so, it is evident that fossiliferous beds now deposited on the shores of North America would hereafter be liable to be classed with somewhat older European beds. Nevertheless, looking to a remotely future epoch, there can be little doubt that all the more modern MARINE formations, namely, the upper pliocene, the pleistocene and strictly modern beds of Europe, North and South America, and Australia, from containing fossil remains in some degree allied, and from not including those forms which are found only in the older underlying deposits, would be correctly ranked as simultaneous in a geological sense.

The fact of the forms of life changing simultaneously in the above large sense, at distant parts of the world, has greatly struck those admirable observers, MM. de Verneuil and d'Archiac. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe, they add, "If struck by this strange sequence, we turn our attention to North America, and there discover a series of analogous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom." M. Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants.

This great fact of the parallel succession of the forms of life throughout the world, is explicable on the theory of natural selection. New species are formed by having some advantage over older forms; and the forms, which are already dominant, or have some advantage over the other forms in their own country, give birth to the greatest number of new varieties or incipient species. We have distinct evidence on this head, in the plants which are dominant, that is, which are commonest and most widely diffused, producing the greatest number of new varieties. It is also natural that the dominant, varying and far-spreading species, which have already invaded, to a certain extent, the territories of other species, should be those which would have the best chance of spreading still further, and of giving rise in new countries to other new varieties and species. The process of diffusion would often be very slow, depending on climatal and geographical changes, on strange accidents, and on the gradual acclimatization of new species to the various climates through which they might have to pass, but in the course of time the dominant forms would generally succeed in spreading and would ultimately prevail. The diffusion would, it is probable, be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. We might therefore expect to find, as we do find, a less strict degree of parallelism in the succession of the productions of the land than with those of the sea.

Thus, as it seems to me, the parallel, and, taken in a large sense, simultaneous, succession of the same forms of life throughout the world, accords well with the principle of new species having been formed by dominant species spreading widely and varying; the new species thus produced being themselves dominant, owing to their having had some advantage over their already dominant parents, as well as over other species; and again spreading, varying, and producing new forms. The old forms which are beaten and which yield their places to the new and victorious forms, will generally be allied in groups, from inheriting some inferiority in common; and, therefore, as new and improved groups spread throughout the world, old groups disappear from the world; and the succession of forms everywhere tends to correspond both in their first appearance and final disappearance.

There is one other remark connected with this subject worth making. I have given my reasons for believing that most of our great formations, rich in fossils, were deposited during periods of subsidence; and that blank intervals of vast duration, as far as fossils are concerned, occurred during the periods when the bed of the sea was either stationary or rising, and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction, and that there was much migration from other parts of the world. As we have reason to believe that large areas are affected by the same movement, it is probable that strictly contemporaneous formations have often been accumulated over very wide spaces in the same quarter of the world; but we are very far from having any right to conclude that this has invariably been the case, and that large areas have invariably been affected by the same movements. When two formations have been deposited in two regions during nearly, but not exactly, the same period, we should find in both, from the causes explained in the foregoing paragraphs, the same general succession in the forms of life; but the species would not exactly correspond; for there will have been a little more time in the one region than in the other for modification, extinction, and immigration.

I suspect that cases of this nature occur in Europe. Mr. Prestwich, in his admirable Memoirs on the eocene deposits of England and France, is able to draw a close general parallelism between the successive stages in the two countries; but when he compares certain stages in England with those in France, although he finds in both a curious accordance in the numbers of the species belonging to the same genera, yet the species themselves differ in a manner very difficult to account for considering the proximity of the two areas, unless, indeed, it be assumed that an isthmus separated two seas inhabited by distinct, but contemporaneous faunas. Lyell has made similar observations on some of the later tertiary formations. Barrande, also, shows that there is a striking general parallelism in the successive Silurian deposits of Bohemia and Scandinavia; nevertheless he finds a surprising amount of difference in the species. If the several formations in these regions have not been deposited during the same exact periods – a formation in one region often corresponding with a blank interval in the other – and if in both regions the species have gone on slowly changing during the accumulation of the several formations and during the long intervals of time between them; in this case the several formations in the two regions could be arranged in the same order, in accordance with the general succession of the forms of life, and the order would falsely appear to be strictly parallel; nevertheless the species would not all be the same in the apparently corresponding stages in the two regions.

ON THE AFFINITIES OF EXTINCT SPECIES TO EACH OTHER, AND TO LIVING FORMS.

Let us now look to the mutual affinities of extinct and living species. All fall into a few grand classes; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, extinct species can all be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the intervals between existing genera, families, and orders, is certainly true; but as this statement has often been ignored or even denied, it may be well to make some remarks on this subject, and to give some instances. If we confine our attention either to the living or to the extinct species of the same class, the series is far less perfect than if we combine both into one general system. In the writings of Professor Owen we continually meet with the expression of generalised forms, as applied to extinct animals; and in the writings of Agassiz, of prophetic or synthetic types; and these terms imply that such forms are, in fact, intermediate or connecting links. Another distinguished palaeontologist, M. Gaudry, has shown in the most striking manner that many of the fossil mammals discovered by him in Attica serve to break down the intervals between existing genera. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals; but so many fossil links have been disentombed that Owen has had to alter the whole classification, and has placed certain Pachyderms in the same sub-order with ruminants; for example, he dissolves by gradations the apparently wide interval between the pig and the camel. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions; but the Macrauchenia of South America connects to a certain extent these two grand divisions. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. What a wonderful connecting link in the chain of mammals is the Typotherium from South America, as the name given to it by Professor Gervais expresses, and which cannot be placed in any existing order. The Sirenia form a very distinct group of the mammals, and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs, without even a rudiment being left; but the extinct Halitherium had, according to Professor Flower, an ossified thigh-bone "articulated to a well-defined acetabulum in the pelvis," and it thus makes some approach to ordinary hoofed quadrupeds, to which the Sirenia are in other respects allied. The cetaceans or whales are widely different from all other mammals, but the tertiary Zeuglodon and Squalodon, which have been placed by some naturalists in an order by themselves, are considered by Professor Huxley to be undoubtedly cetaceans, "and to constitute connecting links with the aquatic carnivora."

Even the wide interval between birds and reptiles has been shown by the naturalist just quoted to be partially bridged over in the most unexpected manner, on the one hand, by the ostrich and extinct Archeopteryx, and on the other hand by the Compsognathus, one of the Dinosaurians – that group which includes the most gigantic of all terrestrial reptiles. Turning to the Invertebrata, Barrande asserts, a higher authority could not be named, that he is every day taught that, although palaeozoic animals can certainly be classed under existing groups, yet that at this ancient period the groups were not so distinctly separated from each other as they now are.

Some writers have objected to any extinct species, or group of species, being considered as intermediate between any two living species, or groups of species. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms or groups, the objection is probably valid. But in a natural classification many fossil species certainly stand between living species, and some extinct genera between living genera, even between genera belonging to distinct families. The most common case, especially with respect to very distinct groups, such as fish and reptiles, seems to be that, supposing them to be distinguished at the present day by a score of characters, the ancient members are separated by a somewhat lesser number of characters, so that the two groups formerly made a somewhat nearer approach to each other than they now do.

It is a common belief that the more ancient a form is, by so much the more it tends to connect by some of its characters groups now widely separated from each other. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages; and it would be difficult to prove the truth of the proposition, for every now and then even a living animal, as the Lepidosiren, is discovered having affinities directed towards very distinct groups. Yet if we compare the older Reptiles and Batrachians, the older Fish, the older Cephalopods, and the eocene Mammals, with the recent members of the same classes, we must admit that there is truth in the remark.

Let us see how far these several facts and inferences accord with the theory of descent with modification. As the subject is somewhat complex, I must request the reader to turn to the diagram in the fourth chapter. We may suppose that the numbered letters in italics represent genera, and the dotted lines diverging from them the species in each genus. The diagram is much too simple, too few genera and too few species being given, but this is unimportant for us. The horizontal lines may represent successive geological formations, and all the forms beneath the uppermost line may be considered as extinct. The three existing genera, a14, q14, p14, will form a small family; b14 and f14, a closely allied family or subfamily; and o14, i14, m14, a third family. These three families, together with the many extinct genera on the several lines of descent diverging from the parent form (A) will form an order; for all will have inherited something in common from their ancient progenitor. On the principle of the continued tendency to divergence of character, which was formerly illustrated by this diagram, the more recent any form is the more it will generally differ from its ancient progenitor. Hence, we can understand the rule that the most ancient fossils differ most from existing forms. We must not, however, assume that divergence of character is a necessary contingency; it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. Therefore it is quite possible, as we have seen in the case of some Silurian forms, that a species might go on being slightly modified in relation to its slightly altered conditions of life, and yet retain throughout a vast period the same general characteristics. This is represented in the diagram by the letter F14.

All the many forms, extinct and recent, descended from (A), make, as before remarked, one order; and this order, from the continued effects of extinction and divergence of character, has become divided into several sub-families and families, some of which are supposed to have perished at different periods, and some to have endured to the present day.

By looking at the diagram we can see that if many of the extinct forms supposed to be embedded in the successive formations, were discovered at several points low down in the series, the three existing families on the uppermost line would be rendered less distinct from each other. If, for instance, the genera a1, a5, a10, f8, m3, m6, m9, were disinterred, these three families would be so closely linked together that they probably would have to be united into one great family, in nearly the same manner as has occurred with ruminants and certain pachyderms. Yet he who objected to consider as intermediate the extinct genera, which thus link together the living genera of three families, would be partly justified, for they are intermediate, not directly, but only by a long and circuitous course through many widely different forms. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations – for instance, above No. VI. – but none from beneath this line, then only two of the families (those on the left hand a14, etc., and b14, etc.) would have to be united into one; and there would remain two families which would be less distinct from each other than they were before the discovery of the fossils. So again, if the three families formed of eight genera (a14 to m14), on the uppermost line, be supposed to differ from each other by half-a-dozen important characters, then the families which existed at a period marked VI would certainly have differed from each other by a less number of characters; for they would at this early stage of descent have diverged in a less degree from their common progenitor. Thus it comes that ancient and extinct genera are often in a greater or less degree intermediate in character between their modified descendants, or between their collateral relations.

Under nature the process will be far more complicated than is represented in the diagram; for the groups will have been more numerous; they will have endured for extremely unequal lengths of time, and will have been modified in various degrees. As we possess only the last volume of the geological record, and that in a very broken condition, we have no right to expect, except in rare cases, to fill up the wide intervals in the natural system, and thus to unite distinct families or orders. All that we have a right to expect is, that those groups which have, within known geological periods, undergone much modification, should in the older formations make some slight approach to each other; so that the older members should differ less from each other in some of their characters than do the existing members of the same groups; and this by the concurrent evidence of our best palaeontologists is frequently the case.

Thus, on the theory of descent with modification, the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms, are explained in a satisfactory manner. And they are wholly inexplicable on any other view.

On this same theory, it is evident that the fauna during any one great period in the earth's history will be intermediate in general character between that which preceded and that which succeeded it. Thus the species which lived at the sixth great stage of descent in the diagram are the modified offspring of those which lived at the fifth stage, and are the parents of those which became still more modified at the seventh stage; hence they could hardly fail to be nearly intermediate in character between the forms of life above and below. We must, however, allow for the entire extinction of some preceding forms, and in any one region for the immigration of new forms from other regions, and for a large amount of modification during the long and blank intervals between the successive formations. Subject to these allowances, the fauna of each geological period undoubtedly is intermediate in character, between the preceding and succeeding faunas. I need give only one instance, namely, the manner in which the fossils of the Devonian system, when this system was first discovered, were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous and underlying Silurian systems. But each fauna is not necessarily exactly intermediate, as unequal intervals of time have elapsed between consecutive formations.

It is no real objection to the truth of the statement that the fauna of each period as a whole is nearly intermediate in character between the preceding and succeeding faunas, that certain genera offer exceptions to the rule. For instance, the species of mastodons and elephants, when arranged by Dr. Falconer in two series – in the first place according to their mutual affinities, and in the second place according to their periods of existence – do not accord in arrangement. The species extreme in character are not the oldest or the most recent; nor are those which are intermediate in character, intermediate in age. But supposing for an instant, in this and other such cases, that the record of the first appearance and disappearance of the species was complete, which is far from the case, we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time. A very ancient form may occasionally have lasted much longer than a form elsewhere subsequently produced, especially in the case of terrestrial productions inhabiting separated districts. To compare small things with great; if the principal living and extinct races of the domestic pigeon were arranged in serial affinity, this arrangement would not closely accord with the order in time of their production, and even less with the order of their disappearance; for the parent rock-pigeon still lives; and many varieties between the rock-pigeon and the carrier have become extinct; and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers, which are at the opposite end of the series in this respect.